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Boraginaceae) Endemic to Sardinia (Italy

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Boraginaceae) Endemic to Sardinia (Italy Systematics and Biodiversity 6 (2): 161–174 Issued 6 June 2008 doi:10.1017/S1477200008002673 Printed in the United Kingdom C The Natural History Museum G. Bacchetta1 , A. Coppi2 , ∗ C. Pontecorvo1 &F.Selvi2 Systematics, phylogenetic relationships 1Centro Conservazione Biodiversit`a (CCB), and conservation of the taxa of Anchusa Dipartimento Scienze Botaniche, University of (Boraginaceae) endemic to Sardinia (Italy) Cagliari, Viale Sant’Ignazio da Laconi, 13 – I09123, Cagliari, Italy Emails: [email protected]; [email protected] Abstract Sardinia is a major centre of diversity of Anchusa, but the systematics, 2Dipartimento di Biologia phylogenetic relationships and conservation status of the taxa endemic to the island Vegetale, Sezione Botanica Sistematica, University of are still poorly known mainly because of their remarkable rarity. We present a Firenze, Via La Pira, 4 – I 50121, study on these endemics based on the results of field research focusing on the Firenze, Italy distribution, the number and size of the populations, the characteristics of the habitat submitted January 2006 and the factors of threat. Combined with observations on aspects of reproductive accepted September 2006 biology, on wild populations and cultivated plants, we evaluate the conservation status of the taxa and propose Red List IUCN categories of vulnerability. Original descriptions, nomenclatural types and karyological data are added. The following Anchusa taxa are endemic to Sardinia: A. capellii, A. crispa ssp. crispa, A. crispa ssp. maritima, A. formosa, A. littorea, A. sardoa and A. montelinasana sp. nov. The latter is described, based on the discovery of a morphologically distinct entity on a mountain in southwest Sardinia. Anchusa littorea was found at a single site after c. 25 years from the last record and 7 years of unsuccessful field research; this species is at the brink of extinction due to the extremely reduced size of the only remaining population. Our complete collection of taxa allowed an analysis of phylogenetic relationships based on DNA sequences from the ITS1 ribosomal genome. In spite of the low variation, this marker produced Maximum Parsimony and Neighbour-Joining phylograms suggesting that the group is monophyletic and that the split between the two clades of the mountain and coastal endemics has been a key evolutionary event. We assume the three mountain species to be relict schizoendemics ancestral to the coastal taxa, and the Paleozoic siliceous massifs of central and south Sardinia as the centre of origin of the group. Key words Anchusa, Boraginaceae, conservation, endemism, molecular phylogeny, Sardinia, systematics Introduction lation size, which explains the lack of herbarium material in many European collections and, consequently, the uncertain Anchusa L. is one of the major genera of tribe Boragineae and often discordant taxonomic interpretations in past and re- (Boraginaceae), with approximately 30 species mainly dis- cent standard floras (Fiori, 1926; Chater, 1972; Pignatti, 1982; tributed in the Mediterranean basin and Middle East, and Greuter et al., 1984). Perhaps the only exception is represented three disjunct African members in the highlands of Eritrea by the psammophytic species, A. crispa Viv., which is the only and Cape region (Gus¸uleac, 1929). Sardinia is a major centre member of the group also occurring on the nearby island of of diversity and endemism for this genus, with six allopatric Corsica. The narrow range, the small size of the populations, taxa occurring in either coastal or mountain habitats of the the loss of habitat caused by human activities and the negative island. However, despite some previous taxonomic contribu- effects of other biotic and abiotic factors are the main causes for tions (Valsecchi, 1976; Selvi, 1998; Selvi & Bigazzi, 1998), which this species is listed in the French and Italian National these narrow-ranged endemics are still poorly known in terms Red Lists (Olivier et al., 1995; Conti et al., 1997; Scoppola & of both phylogenetic relationships and conservation status. A Spampinato, 2005), in the EU Directive 92/43 CEE ‘Habitat’, reason for this is their very restricted range and small popu- in the Berne Convention and finally in a recent IUCN selec- tion of the 50 most endangered species of the Mediterranean ∗Corresponding author. Email: selvi@unifi.it (De Montmollin & Strahm, 2005). Accordingly, A. crispa has 161 162 G. Bacchetta et al. been the object of numerous studies on different ecological number of sites recorded previously for each taxon on the and reproductive aspects related to the problems of its in situ basis of reliable literature data and herbarium specimens in FI, conservation, though almost exclusively on Corsica (Thiebaut,` CAG, SASSA, TO and G. For the rarest species we prepared 1988; Guyot & Muracciole, 1995; Paradis & Piazza, 1988, fine scale maps, which will allow the detection of eventual 2000; Quilichini & Debussche, 2000; Quilichini et al., 2001, future changes in the area of occupancy of the populations. 2004). Based on these data, we propose conservation categories for It is evident, therefore, that the disparity of knowledge each taxon following the IUCN criteria (IUCN, 2001). In ad- that we have for this ‘target’ species with respect to the other dition, collection of seed allowed us to make preliminary tests Sardinian endemics even in basic terms of number and surface on germination capacity and to grow plants in the botanical of the populations is not a favourable condition to plan initiatit- garden for the purposes of ex situ conservation. During the ives for their long-term, in-situ conservation. To fill this gap we years 2000–2005 we monitored the production of viable seed undertook a field-based research to obtain more information in isolated plants under cultivation. on the distribution, habitat features and demographic condi- Before treating the above-mentioned aspects, we give for tion of the populations. Seven years of field work allowed us each taxon: (1) the nomenclatural type; (2) a brief morpholo- to find new localities for some of the rarest taxa, to better gical description based on field notes and herbarium material; understand the biotic and/or abiotic factors that influence the (3) references to selected iconographies and an original illus- demographic dynamics of populations, and to establish cor- tration of the new species A. montelinasana. The latter species rect IUCN categories. This is the essential information to plan and the new population of A. formosa from Mount Arcosu recovery actions of all endangered species (Cropper, 1993; (see below) were also analysed from a karyological viewpoint, Frankel et al., 1995). In addition, we discovered a popula- using root tips of seeds germinated in Petri dishes. These were tion of Anchusa in a rugged mountain area of the southwest- pre-treated with 0.002 M 8-hydroxyquinoline or 0.05% col- ern part of the island, whose morphological distinctiveness chicine, 2.5 h at room temperature and then fixed overnight allowed us to describe it as a new species. Such an import- in ethanol:glacial acetic acid 3:1. The meristematic tissue was ant discovery made possible a complete collection of mater- then thoroughly rinsed in distilled water, hydrolysed in 1N ial for the phylogenetic analysis of the group using sequences HCl at 60 ◦C for 6–7 minutes, and stained in lacto-propionic from the non-coding Internal Transcribed Spacer region (ITS1) orcein overnight. Meristems were finally dissected, squashed of the nuclear ribosomal DNA. The usefulness of this molecu- on clean glass slides in a drop of 45% acetic acid, and observed lar marker in the species-level systematics of Boragineae has with a Zeiss Axioscop light microscopy under oil immersion been demonstrated in recent studies (Hilger et al., 2004; Selvi (100×). et al., 2004). The results of this work are reported in this paper in order to contribute to a better knowledge and to the con- DNA isolation and amplification servation of this remarkable component of the Mediterranean Genomic DNA of the taxa listed in Table 1 was extracted × insular endemic flora. following a modified 2 CTAB protocol (Doyle & Doyle, 1990) using silica-gel samples of leaf tissue collected in the field by the authors. Voucher specimens are deposited in FI, Materials and methods CAG and BM. The extracted DNA was quantified after agarose gel electrophoresis (0.6% w/v) in TAE buffer (1 mM EDTA, Fieldwork and plant material 40 mM Tris-acetate) containing 1 μg/ml of ethidium bromide This account is mainly based on fieldwork (1998–2005) fo- by comparison with a known mass standard. The primers ITS4 cusing on the distribution, ecology and conservation status of and ITS5 (Baldwin, 1992) were used for the amplification of the taxa in terms of number and area of occupancy of the the Internal Transcribed Spacer ITS1 region of nuclear rDNA populations, approximate number of fertile individuals, hab- intron. PCR amplifications were performed in a total volume itat characteristics and main factors of threat. When possible, of 50 μl containing 5 μl of reaction buffer (Dynazyme II; we captured the insects that visited the flowers and made ob- Finnzyme, Espoo, Finland), 1.5 mM MgCl2, 20 pmol of each servations on the means of seed dispersal. We determined the primer, 200 μM of each dNTP, 1 U of Taq DNA polymerase Taxa Geographic origin and herbarium vouchers A. capellii Moris Sardinia, Esterzili, Monte S. Vittoria, G. Bacchetta & F. Selvi, 99.002 (FI) A. crispa Viv. ssp. crispa Corsica, Solenzara, Cannella, M. Bigazzi & F. Selvi, 99.005 (FI) A. crispa Viv. ssp. crispa Sardinia, Alghero, Porticciolo, M. Bigazzi & F. Selvi, 97.001 (FI) ““ ssp. maritima (Vals.) Selvi & Bigazzi Sardinia, Badesi, mouth of river Coghinas, M. Bigazzi & F. Selvi, 97.005 (FI) A. formosa Selvi, Bigazzi & Bacchetta Sardinia, Uta, Mt. Lattias, M. Bigazzi & F. Selvi, 97.006 (FI, CAG) A. littorea Moris Sardinia, Arbus, Is Arenas, G. Bacchetta & C. Pontecorvo (CAG) A. montelinasana Sardinia, Gonnosfanadiga, Mt.
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